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Alien Flora in Grasslands Adjacent to Road and Trail Corridors in Glacier National Park, Montana (U.S.A.) Author(s): Robin W. Tyser and Christopher A. Worley Source: Conservation Biology, Vol. 6, No. 2 (Jun., 1992), pp. 253-262 Published by: Blackwell Publishing for Society for Conservation Biology Stable URL: http://www.jstor.org/stable/2386247 . Accessed: 09/09/2011 11:20
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AlienFlorain Grasslands to Adjacent Road and Trail in National Corridors Glacier (U.S.A.) Park,Montana
ROBIN W. TYSER
Department of Biology and Microbiology University of Wisconsin-La Crosse La Crosse, WI 54601, U.S.A.

CHRISTOPHER A. WORLEY
Department of Biology and Microbiology University of Wisconsin-La Crosse La Crosse, WI 54601, U.S.A.

Abstract:Alien plant species have rapidly invaded and successfullydisplaced native species in many grasslands of NorthAmerica Thus, the status of alien species in western thenature reserve grasslands of thisregionwarrantsspecial attention. This study describes alien flora in nine fescue grassland studysites adjacent to threetypesof transportation corridors-primaryroads, secondary roads, and backcountrytrails-in Glacier National Park,Montana (USA). Parallel transects, placed at varyingdistancesfrom the adjacent road or trail, were used to determinealien species richness and frequency at individual study sites. Fifteen alien species were recorded,two Eurasian grasses, Phleum being particularly common in pratense and Poa pratensis, most of the study sites. In sites adjacent to primary and secondary roads, alien species richnessdeclined out to the most distant transect, suggestingthatalien species are successfullyinvading grasslands from the roadside areas. In trails,absence of a comstudysitesadjacent to backcountry highlevelsof alien species parable decline and unexpectedly richness100 m from the trailsidesuggestthatalien species have been introducedin off-trail areas. The resultsof this study imply that in spite of low levels of livestockgrazing and otheranthropogenicdisturbances, fescue grasslands in nature reserves this regionare vulnerable to invasion by of alien flora Given theprominentrole thatroadsidesplay in theestablishment and dispersalofalien flora road construction should be viewed from a biological, rather than an
Paper submittedSeptember25, 1990; revisedmanuscriptaccepted March 28, 1991.

Resumen: Especies de plantas introducidas ban invadido rapidamentey desplazado exitosamenteespecies nativas en praderas del Oeste de America del Norte.Por lo tanto el estado de las especiesintroducidasen las reservas depastizales naturalesde esta regionexige especial atencion.Este estudio describela flora introducidaen nueve pastizales naturales de defestuca las acreas estudiosson adyacentesa trestiposde corredoresde transporte-caminos primarios, caminos secundariosy senderosremotos- en el Parque Nacional "Gla" riqueza y frecuencier, Montana (EE. UU). Para determinar cia de especies introducidas, se trazaron transectas paralelas, localizadas a distancias variables del camino o sendero adyacente en las areas de estudio. Se registraron quince especies introducidas. Dos pastos eurasiacticos, Phleumpratensis Poa pratensis, y resultaron particularmente abuntes en la mayoria de las acreasde estudio. En lugares adyacentesa caminosprimariosy secundarios,la riqueza de especies introducidas disminuyo en la direccion de las transectasma's distantes,sugiriendo que las especies introducidas estan invadiendo exit6samentelas praderas desde areas aledanias a caminos. En las acreasde estudio adyacentesa senderosremotosno se encontr6una disminucion comparable; inesperadosaltos nivelesde riqueza de especies introducidasa 100 m de los senderos,sugieren que las especies foracneasban sido introducidas desde otras areas Los resultadosde esteestudio implican fuerode los senderos. que a pesar de los bajos niveles de pastoreo y otrasperturbaciones antropogenicas, los pastizales de festuca en las reservasnaturales de esta regionson vulnerablesa la invasion de laflora introducida Dado el rolpreponderanteque 253
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AlienFloraAdjacent Road and TrailCorridors to

Tyser Worley &

engineeringperspective.Nature reserveman agers should establish effective roadside vegetation management programs that include monitoring quickly treatingkeystone alien species upon theirinitial occurrence naturereserves, in and creating buffer zones on roadsides leading to nature reserves.

juegan los caminos en el establecimiento dispersionde la y flora introducida la construccionde rutas debe ser vista desde un punto de vista biol6gico, mas que desde una perspectivameramente ingenieril. administradores reserLos de vas naturales deberi'an establecerprogramas efectivosde maneijo de vegetacion en los bordes de los caminos. Estos programas deberian incluir monitoreo,tratamientorapido de especiesintroducidasyclaves tanpronto como se detecten en las reservasnaturales,y creacion de zonas de transicion en los caminos que conducen a las reservasnaturales.

Introduction
The widespreaddisplacement nativeplantspecies by of alien florain the Intermountain bunchgrasscommunities ofwesternNorthAmericahas been consideredin a numberof studies(Young et al. 1972; Mack & Thompson 1982; Mooney et al. 1986; Mack 1986). Mack (1989) notes that this displacement is essentiallya worldwide phenomenon,occurringin the bunchgrass communitiesof California'sCentral Valley, southern SouthAmerica, and southern Australia. Successfulalien plants in the Intermountain grasslands-grasslandsin the regionbetween the RockyMountainsand Cascade Mountains-include species considered agriculturally useful(e.g.,Poapratensis) as well as species considered noxious (e.g., Centaurea maculosa-Watson & Renney 1974). Baker(1986) and Forcellaand Harvey (1983) suggest that high lightintensity and frequentbreaks in plant cover characteristic grasslands of invasion mayfacilitate by alien species. In addition,the success of alien vegetationin grasslands regularly is attributed anthropoto genic disturbancessuch as cultivation, to attempts reduce shrubcover,and especiallygrazingand trampling by domesticungulates(Daubenmire 1970; Young et al. 1972; Mack 1981; Mack & Thompson 1982; Mooney et al. 1986). Alienspecies are knownto possess a variety of adaptationsto disturbance, includingrapid refoliation and photosynthetic recovery,prostrategrowthform and ground-level and buds,highlevels of sclerenchyma lignified cells, and phenotypic leaf plasticity (Shearman & Beard 1975; Caldwell et al. 1981; Bazzaz 1986; Hall & Kuss 1989; Butt et al. 1989). Significantly, Rejmanek's (1989) simulationmodels suggestthat even competitivelyinferior alien species can successfully invade nativecommunities, givenappropriately highlevels ofdisturbance. Though nature reserves are presumablyrelatively protected from the effectsof human-relateddisturbance, a numberof alien species have been recordedin grasslands nationalparksalong the easternborderof of the Intermountain Glacier (U.S.A.), area, thatis, Banif, Jasper,Grand Teton, and WatertonLakes (Koterba &

Habeck 1971; Stringer 1973; Weaver & Woods 1985, 1986). Because roadsides are human-disturbed microhabitatsto which alien plants are oftenwell adapted (Frenkel1970; Forcella& Harvey1983), it is reasonable to expect the occurrenceof alien florain nationalpark grasslands be correlatedwith adjacent roadside corto ridors.Trailsideareas are also human-disturbed corridors (Dale & Weaver 1974; Bright1986; Hall & Kuss 1989) ofpossible importance the occurrenceofalien to species in these grasslands. As theinitial phase ofa long termmonitoring project, the present study considers the alien flora of fescue grasslands adjacent to primary roads, secondaryroads, and backcountry in trails GlacierNationalPark, Montana (U.S.A.). Dominantnativegrassesin thishabitatinclude Festuca idahoensis, F. scabrella, and several other bunchgrassspecies. This studyaddresses two specific questions:(1) Whatalien species have mostsuccessfully invaded grasslands adjacent to road and trailcorridors in thepark?and (2) To what extentcan thisinvasionbe attributed the effects road and trailcorridors? to of

Sites Descriptionof Study


Nine fescue grassland studysites in fivedifferent drainage systems Glacier NationalPark,Montana(U.S.A.), of were selected for study.Total grasslandarea in these drainageswas ca. 2800 ha. Vegetationwas representative of theFestuca scabrellaiF. idahoensis habitattype of Muegglerand Stewart(1980). In addition to these two species, other native grasses common at various study siteswereAgropyron caninum,Koeleria cristata Danthonia intewnedia and Stipa richardsonii.Common forbsincludedAcbillea millefolium,Galium boreale, Cerastiumarvense,and Lupinus sericeus (Tyser 1991). A cryptogam ground layer composed of small lichens and mosses coveringundisturbedsoil surfaces was commonly presentin these grasslands. Primary roads in Glacier Park are narrow,asphaltbased, two-laneroads occasionallyresurfaced with oil and rock chip treatments. Roadside shouldersare usually mowed once or twice each year. Secondaryroads are unimproved dirtroads bladed by road maintenance

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equipmentabout once per year,but otherwisetheyreceive littlemaintenance.Backcountry trailstudysites or were >2 km fromthe nearest primary secondary road. Only hikingand horse use are permitted park on trails. Studysites were relatively topographically homogeneous, >1 km fromother studysites,and bordered a primary road, a secondaryroad, or a backcountry trail segmentwith less than a 100 curve. Of the nine study sites, three (PR1, PR2, PR3) bordered primary roads, three (SRI, SR2, SR3) bordered secondaryroads, and three (BT1, BT2, BT3) bordered backcountrytrails. Studysite locations are indicatedin Figure 1. Withthe exception of sporadic homesteadingthat occurred in the northwest section of the parknear the SR1 and SR2 study sites and a short-lived miningoperation abandoned ca. 1900 near the BT1 site (Fig. 1), no records existindicating of thatgrasslands theparkwere used for livestockor agricultural activitiesbeforethe park was established in 1910. Mean annual precipitationand mean dailyJulytemperatures fromrecordingstations closest to the studyareas are SRI and SR2-59.2 cm, 15.40 C; PR1, PR2, and SR3-67.1 cm, 15.80 C; PR3 and BT1-99.1 cm, 13.80 C; BT2 and BT3-96.5 cm, 14.30 C (Finklin1986). Withthe exception of the BT1 site thatwas adjacent to a trail establishedin ca. 1974 (J. Potter,personal all communication), studysiteswere borderedby roads or trailsthathave existed in the park since at least the 1930s (Ruhle 1972). The road adjacentto theSR3 study sitehas been closed to vehicle traffic used as a trail and since the early 1970s, thoughit was a secondaryroad from1930 to 1970. Because of its longeruse as a road, it was classified a secondaryroad studysite. as
GlacierNational Park

Methods
Study Design Studysiteswere sampled duringa 5-weekperiod from lateJuneto earlyAugustin 1988 and 1989. Species that could not be reliablysampled throughout this 5-week interval, example springephemerals, for were not considered.Withone exception(see below), studysite dimensionswere 100 m X 100 m. In the six road study sites a 100 m transect, designatedas the "R" transect, was placed in the centerof the roadside area between theroadbedand thegrassland border(Fig. 2). Typically, roadside areas consisted of a drainage ditch (primary roads) or a ridgeof substrate pushed to the side of the road bed (secondaryroads). In all nine studysites, three 100 m transectswere placed at varying distancesfromthe grasslandborder parallel to the road or trail(Fig. 2). The first transect, as designated the "G1" transect, placed 1-2 m from was thegrassland border.Two othertransects, designatedas
Road Roadside

Road Primary and Road Secondary StudySites

100m

01

02

03

BT2

Trail Backcountry StudySites

25m 25 mm

SR 1~~~~B1 *SRI

~~~~~~~BT3
PR3

Monan

6SR2~~~~~~R
N
W E

Trail

Pi SR3
Primary Road Number Study of Sites Site Study Acronyms 3 PR3 PRI,PR2,

Transect Corridor Types

Grassland

5 10 0 kilometers

Continental Divide

Secondary Road 3 SR3 SRt,SR2, R, G1,G2,G3

Backcountry Trail 3 BT1, BT2,BT3 G1,G2,G3

Road Primary Road Secondary Site * Study ParkBoundary

Site R, G1,G2,G3 InEachSudy Transects

Figure 1. Map of Glacier National Park showing studysite locations.

Figure2. Location of theA GI, G2, and G3 transects forprimaryroad, secondaryroad, and backcountry trail studysites.Note that transects comparable to roadside ("R") transects were not used trail sites. for sampling the backcountry

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"G2" and "G3," were located 25 m and 100 m, respectively, fromthe Gl transect.Because of encroaching forestvegetationat the BT3 site, the G3 transect was positioned 75 m ratherthan 100 m fromthe grassland edge. Thus, alien vegetationwas sampled along four transects Gl, G2, G3) in each of the six road study (R, sites and along threetransects (Gl, G2, G3) in each of the three backcountry trail study sites. Roadside transects were in areas clearlydisturbed road-related by activities such as mowing,blading,road repair,and vehicle traffic. However,otherthanoccasional depositsof soil and smallrocksalongseveraloftheGl transects, no indications human-related of substrate disturbance were apparentalong any of the G1-G3 transects. Presence of all alien vascularspecies was determined in 20 cm X 50 cm quadratsat 2 m intervals along each transect.Percentage frequencyof individual species (percentage of quadrats in which the species was observed) and mean alien species richness(mean number of alien species per quadrat) were determined each for transect.Use of alien species richnessis a simple yet in effective way to detect differences vegetationpatin ternsamongtransects these grasslands (Tyser & Key and 1988). Species nomenclature determination alien of status followHitchcockand Cronquist(1973). It is possible thatPoa pratensis, or a morphologically similar species, is nativein remote areas in the Intermountain region of western North America (Cronquist et al. 1977). In this study,P. pratensis was considered an alien species. Data Analysis The nine studysites were divided into three corridor road sites,secondary road sites,and backtypes: primary trailsites. The unit used forstatistical country analysis was the number of alien species per 0.1 m2 quadrat, designatedas alien species richness.For each corridor of type,the effects transectand site on alien species richnesswere evaluated using two-wayANOVAs followed by post hoc Tukey multiplecomparisons.Because alien species richnessdata fromseveraltransects and could not be transwere not normally distributed or formed standard transforby logarithmic square-root alien species richnesses were rank-transformed mations, for each corridor type prior to ANOVA. Two-way ANOVA of ranksis a robust,nonparametric approach describedby Conover and Iman ( 1981 ) and references therein.SYSTATv. 3.2 software (Wilkinson1987) was used to perform statistical computations.

teraceae (four species), Fabaceae (three species), and Poaceae (fourspecies). Eightofthesespecies were sampled along the G3 transects thatwere located farthest from possible effects adjacentroad and trailactivities. of Three additionalobservationsare worth notingabout species occurrencesassociatedwith the threecorridor types.First, severalspecies-most notably Pbleum pratense, Poa pratensis,and to a lesser extentTaraxacum officinale-were commonly recorded (frequency >20% ) along one or more transectswithin all three corridor types. Second,thenumberofalien species sampled along the G1-G3 transects was surprisingly similar among the three corridortypes.For example, the cumulativenumbersof species sampled along the G1-G3 in transects theprimary road studysiteswere three, six, and fourspecies per site, respectively. Corresponding numbersfor the other two corridortypeswere four, and six species per studysite forsecondaryroads four, and three, four, and fivespecies per studysite forbackcountry trails. Third,the occurrenceof Centaurea maculosa was consistently associated with the primary road studysites.Centaurea maculosa frequencies were relatively highalong the R and Gl transects primary in road studysites,but this species was not recorded in any of the secondaryand backcountry trailstudysites. Thiswas a typical pattern observedthroughout park. the Continuity Roadsideand Trailside with Areas Most occurrence patternsof individualspecies among transects withinstudysites were continuouswith the transect closest to the adjacent road or trail(Table 1).
sampling pattern,

forexample,Melilotus officinalisat the PR1 studysite, where a species was recorded along the R transect but not recorded along the Gl, G2, and G3 transects. Anmaculosa at the PR2 studysite,where a species was recorded along the R and Gl transects, not the G2 but or G3 transects. Thoughsome species exhibitedgaps in theirdistributions outwardfromthe road or trail(patterns such as "- + - + " or " + - + -," e.g., Poa compressa at the SR2 site), these instanceswere relatively few.Of the 46 species occurrencepatterns, onlyninetwo in theprimary road sites,fivein the secondaryroad sites,and two in the backcountry trailsites-exhibited such distribution gaps (Table 1). AlienSpeciesRichness Patterns
ALIENSPECIES RICHNESSGRADIENTS

This was exemplified by a " + ---"

other pattern was " + +

- -,"

for example, Centaurea

Results
SpeciesSampled Fifteenalien plant species were recorded (Table 1), most of which were members of three families:As-

Withinall three corridortypes,alien species richness 1 (number of alien species/0. m2 quadrat) varied significantlyamong transectsand among studysites (Table 2). Alien species richnesstypically declined monotoni-

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Alien Flora Adjacent RoadandTrail to Corridors 257

Table 1. Percent frequency alien speciesbytransect. = 50 quadrats eachtransect. of N for


Site Transect ALY BRO CEN 100 74 2 88 64 18 CIR
-

LAP
-

LIN
-

MEL 54
-

PHL 50 6 4 80 60 74 28 88 34

POC

POP 60 28

PRU
-

TAR

TRA

TRP

TRR

a. Primary Roads R PRI G2 G3 R GI G2 G3 R

GI

PR2

18 8
-

PR3

G2 G3 b. SecondaryRoads R SR1 Gl SR2 G3 R G2 G3 R

GI

6 2

48 50 14 4 58 38 82 26

30 12 6 6

6 2
-

G2

GI

40 18 16 6

18 16 2 2

32 20 22 2 8 78 98 92 98

2 2

100 84 8 62 74 32 16 34 18 20 64 6 2 74 24 12

28
-

2
-

80

SR3

G2 G3 c. Backcountry Trails GI BT1 G2 G3 BT2 Gl G2 G3 Gl BT3 G2 G3

GI

22

6
2

6 8 2

66 12 4 94 68 68 100 90 96

16 6 16 28 24 60

2
14

aSpecies acronyms:ALY (Alyssumalyssoides),BRO (Bromus inermisssp. inermis),CEN (Centaurea maculosa), CIR (Cirsium arvense), LAP MEL (Melilotus officinalis), PHL (Phleum pratense),POC (Poa compressa),POP (Poa pratensis), (Lappula echinata),LIN (Linaria vulgaris), PRU TRA(Tragopogon dubius), TRP (Trifolium (Prunella vulgarisvar. vulgaris),TAR(Taraxacum officinale), pratense),TRR (Trifolium repens).

in callyfrom R to G3 transects theprimary the road and secondary road sites(Table 2a, b; Fig.3). However,alien species richnessin the backcountry trailsites did not show a G1-G3 decline (Fig. 3). Though alien species richnesswas significantly greateralong the Gl transect than along eitherthe G2 or G3 transectsin the backcountry trailsites,the G2 and G3 transects notdiffer did significantly one another(Table 2c). Hence, there from was an alien species richnessgradient the backcounin trystudysites,but it did not extend as deeply into the grassland was the case forthe primary secondary as and road sites.
AMONG CORRIDORTYPE COMPARISONS G1-G3 TRANSECTS OF

grasslandtransects two backcountry in trailsites,particularly BT3 site. the

Discussion
Alien SpeciesRichness Patterns One of the more consistent patterns foundin thisstudy was the R-G3 decline in alien species richnessforprimaryand secondaryroad studysites.Because distributionsof most alien species did not exhibitgaps in their distributions outward fromthe R transects, this alien species richness gradient probablyrepresents more or a less continuousdecline in the frequenciesof individual species. These patternsare consistentwith models of species invasions (Williamson& Brown 1986; Rejmanek 1989; Williamson1989) where an invading species progressively spreads ("diffuses")fromits point of initial introduction, which in thiscase would be the roadside area. Hence, the alien species richnessgradients can be attributed road-related to effects. Some roadside introductions have undoubtedlyre-

Alien species richnessalong the G1-G3 primary road transectswas not consistently higher relative to the othertwo corridortypes.In fact,overall quadratrichness means of the G2 and G3 transects the primary at meansofboth road siteswere lower thancorresponding secondary road and backcountrytrail sites (Fig. 3). Alienspecies richness was relatively highalongall three

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Table 2. Summary two-way of of ANOVAs theeffects transect of location study on alien speciesrichness and site (number alien of m2 species/0.1 quadrat)forthethreecorridor PosthocTukey types. are multiple for comparisons indicated thetransect factor.
Comidor type Factor SS df F p Tukeya

a. PrimaryRoads

b. SecondaryRoads

c. BackcountryTrails

TransectLocation StudySite Transect x Site Error TransectLocation StudySite Transect x Site Error TransectLocation StudySite Transect x Site Error

9549961.09 1654988.16 439738.35 4345810.40 5648871.45 5302817.59 444031.53 4571449.93 1151136.33 2250215.23 133959.37 3285676.57

3 2 6 588 3 2 6 588 2 2 4 441

43.43 111.96 9.92 25.44 341.04 9.52 17.19 151.01 4.50

<0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.011 <0.001 0.001

R > Gl > G2 > G3

R > Gl > G2 > G3

Gl > G3 = G2

a Transectsare listed in decreasingorderof mean alien species richnessranks. Transectsdiffering in significantly alien species richness(p < 0.05, Tukeymultiple comparison tests)are indicated by ">".

suitedfromaccidentaltransport alien seeds by vehiof cles (Schmidt 1989). However,intentional roadsideintroductions are an additional, potentiallyinfluential source of alien species. Untilca. 1980, revegetation efforts parkpersonneland road construction by contractorscommonly includedseedingprimary secondary and roadsides with alien seed mixes consistingof species spp. (D. Lange,personalcommunication). additionto In to contributing the alien species richnessgradient, such seedingsmayhelp explainthe consistent occurrenceof P. pratense andP. pratensis in all studysitesadjacentto primary and secondaryroads. Other factors, disespeciallyhighlevels of substrate turbancein the roadside areas, may help maintain the alien species richnessgradient foundin theprimary and secondary road sites. Substratedisturbanceprobably promoteshigh alien species populationdensitiesalong roadsides,thusproviding abundantseed sourcesforongoingdispersalintoadjacentgrasslands. Such is thecase forinvading Centaurea maculosa colonies,where roadside seed production was estimated be about threeto to fourtimeshigherthanin the adjacent grassland (Tyser & Key 1988). Unliketheprimary secondary and road sites,thebacktrailalien species richnessgradient not excountry did tendpast the G2 transect (Table 2c). Thus,traileffects on alien species distributions were less prominent than road effects. Interestingly, alien species richness G3 was relatively highin two of the trailsites (Fig. 3), suggesting thatintroduction alien species has not been conof finedto trailsides. fact, In unauthorized seedingof alien grassesby horse concession personneland pasturing of concession horses did occur priorto the mid-1940sin backcountry grasslandseast of the continentaldivide (Park Records 1939; J. Potter,personal communication). If off-trail introductions occurred,the shorttrailside alien species richnessgradientcould resultfrom additionaltrailsideintroductions hikersand horses by
such as Pbleum pratense, Poa pratensis, and Trifolium

or from trailside selection(e.g.,from trampling-Dale & Weaver 1974; Cole 1987; Hall & Kuss 1989) fordisturbance-tolerant species initially introducedin off-trail areas. At any rate,an explanationcomparable to thatofferedforsites adjacent to roads-trailside introduction and subsequentdispersalinto adjacent areas-does not account for alien distribution sufficiently patternsobservedin the trailsites. of Comparisons theG1-G3 transects amongthethree corridortypesshowed two unexpected patterns. First, the number of species sampled along the grassland transectswas relativelysimilarfor all three corridor types(see "Species Sampled"in Results).Second,lower alien species richness was expected in the backcountry sites,since these sites are characterizedby lower visitation rates, absence ofroadsideditches, an and presumably lower levels of alien seed introduction than the road sites.However,such anticipateddifferences were notobserved(Fig. 3). Indeed,alien species richness was relatively highat the BT3 site.Likewise, mean alien species richnessalong the G2 and G3 transects the secin ondary road sites was not lower than corresponding means forthe primary road sites. These unexpectedpatterns suggestthatintroduction rateswithintheparkhave notbeen well correlated with typeofroad or trailcorridor. example,past roadside For seedingsby resourcemanagers maynot have been proportionalto road use intensity, unauthorized and introductionsby horse concessionpersonnelmayhave been commonin some backcountry relatively grasslands. is It also likelythatvariations site-specific in for factors, example, compositionof native vegetation,fire history, alien florain ways not associated substrate, etc., affect with the adjacent transportation corridor.Variationin precipitation does not appear to be a site-specific factor that correlateswell with the presence of alien flora, however.For example,alien species richnessalong the G1-G3 transects relatively in thePR3,SR1,SR2, was low and BT1 sites(Fig. 3). Yet the SR1 and SR2 sitesreceive

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259

2.8

A. PrimaryRoad Sites
R

Phleum & pratense

Prominent AlienSpecies Several species, particularly Pbleum pratense and Poa pratensis,were relatively common in most studysites (Fig. 3). In addition,roadside colonies of Centaurea maculosa monitored since 1984 are currently invading adjacent grasslands(Tyser & Key 1988; Tyser 1991). Whatfactors contribute the success of these species? to Mack and Thompson (1982) and otherspropose that because Eurasiangrasseshave had a more extensiveevassociationwith ungulategrazersthan have olutionary westernbunchgrasses, theyare comparatively resistant to grazing and trampling effects. example,following For simulatedgrazing,an alien grass,Agropyrondesertorum, establishesa canopy with threeto fivetimes the photosynthetic surface area ofa morphologically similar native species, Agropyronspicatum (Caldwell et al. 1981). The selectiverole ofdomesticlivestock a comis the of monlycitedfactor underlying replacement native species by alien grassesand forbsin the Intermountain west and elsewhere (Young et al. 1972; Mack 1989). Likewise,the widespread success of noxious forbsin thisregion,such as C maculosa, is typically attributed to overgrazing livestock and other anthropogenic by substrate disturbances (Watson & Renney 1974; Lacey et al. 1986). However,the overalllevel ofdomesticlivestock grazin ing and otherhumandisturbances the parkhas been relatively low, and Mack (1986) notes instanceswhere alienspecies have invadedgrasslands knownto have not been disturbed humans.While domesticgrazing and by anthropogenicdisturbances may hasten invasion by alien species, our results suggest that dispersal from road and trailcorridorsinto adjacent naturalareas can occur even iflevels ofthesefactors low, and perhaps are even if theyare absent. Other factors, includingtolerance to grazingand trampling elk, reduced biotic by controls,and preadaptationto steppe environments, mayaccount forthe success of alien species in natural area grasslandsof this region. Mechanismsunderlying the success of alien species in agro-systems may also For operate in naturalsystems. example,the success of C. maculosa in rangelands maybe increasedby bittertastingcnicin, a sesquiterpenelactone found in high in concentrations thisspecies (Locken & Kelsey 1987). Cnicin may also renderthis species unpalatableto native mammaland insectherbivoresin naturereserves. Aftersuccessfully invadingnaturalarea bunchgrass to communities, what extentdoes alien floraalterhost Additionalobservationsin the park sugcommunities? gest thatPhleum pratense, and especially Centaurea maculosa, are capable ofdecreasingcover and diversity of native species (Tyser 1992; see also Tyser & Key 1988). In addition, cryptogamground cover, which role in soil stabilization, plays an important nitrogen fixation, and water relationships the Intermountain in

2.4 2.0 1.6


1.2

R
01 R

Grasse-s mOther EJForbs

Poa pratensis

01I

02S

0.8 0.4
0 0 02 3 023 023

PR1 B. Secondary Road Sites

PR2
SR

PR3

2.8
2.4-

2.01.6of1.2

0pi

0.8~~~~~~~~~0
02

0.4

-G3

02

0.0critiono 2.8
2.4-

SRt

SR2

SR3

C. BackcountryTrail Sites
01

2.0

GIG
01I

1.6
1.2

02 03 02023 01I G

0.8 0.4 0.0

02

BTI

BT2 Study Sites

BT3

Figure3. Summaryof alien species richness(number m2 of alien specieslO.t1 quadrat) along each transect in thenine studysites.Arrowsindicate overall transectmeans withineach corridortype. littleprecipitation, while the PR3 and BT1 relatively sites receive relatively highlevels of precipitation (see Descriptionof StudySites). Now that horse-relatedbackcountryactivities are more carefullyrestrictedand roadside seeding with alien species has stopped in the park,future introductionratesare expected to be higher roads alongprimary trails. thanalong secondaryroads and backcountry The recentdispersalof Centaurea maculosa into relatively Glacier National Park,a species firstobserved in the parkin the mid-1960s(R. Wassem,personal communiroadcation), and itspresentoccurrencealongprimary to sides probablycharacterizes colonizationpatterns be the followedby future species entering park.ThoughC maculosa and future species are undoubtedlycapable it of colonizing secondaryroadsides and trailsides, is roadoccur along primary likelythattheywill initially corridor sides. Thus, distinctions among transportation years typesshould become more pronouncedin future thantheywere duringthisstudy.

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region(Reddy & Giddens 1975; Andersonet al. 1982; Brotherson Rushforth & 1983), was lower in areas dominated by C. maculosa and P. pratense than in areas dominatedby native graminoids(Tyser 1992). Mack and Thompson(1982) suggestthatrhizome-tiller mats formedby alien grasses restrictcryptogam coverage. Though additionalstudyis clearlyrequired,the potentialeffect alien vegetation nativebunchgrass of on communities a legitimate is concern to naturereservemanagers. In conclusion,our studyshows thatseveral species, now widelyoccur in fescue especiallyEurasiangrasses, grasslands adjacent to road and backcountry trailcorriand thatdispersalofthese dors in GlacierNationalPark, has species fromroadsides into adjacent grasslands octhree issues of general curred.These resultshighlight concern to the managementof nature reserves.First, commonlycited anthropogenicdisturbancessuch as livestockgrazing not sufficiently do account forthe ocIt currence of alien species in these grasslands. seems communities not are clear thatnaturalarea bunchgrass immune from alien species invasions. Second,mostalien introductions probablyoccurredafterthe parkwas established.This underscoresthe need for alien vegetation managementprogramsthat are proactive,rather thanreactive.Third,thoughroads and otherline corriof dors can benefit the migration nativespecies within landscapesfragmented humandisturbance by (Saunders et al. 1991), our observations show thatsuch corridors influence naturalarea grasslands. negatively

Management Implications
are proposed. Several managementrecommendations actions should be careThese and other management and expeditiously fully considered,given the apparent in of vulnerability naturereservegrasslands thisregion to invasionby alien flora. 1. This study provides yet another reason to avoid in road-building naturereserves.Where road constructionin bunchgrasscommunitiesis unavoidable, the originaltopsoil should be redepositedin roadsideditches.In addition, road construction projects should not be considered complete untilnative roadside vegetationis fullyestablished.Propin erly viewed, road construction naturereserves as should be treated, and appropriately funded, 1020-year biological projects, ratherthan 1-2-year engineering projects.Thus,biologistsand resource rather managersshould oversee road construction, thanengineers. 2. Given the success of alien grassesthatwere intenseeded in the park,an obvious recommentionally all introductions of dationis to eliminate intentional alien species, even in developed areas (e.g., road-

sides), withinnaturereserves.In additionto being invasive,alien species used for revegetationhave been shownto suppressestablishment nativespeof cies (Wilson 1989). 3. Because roadsidesare especiallyvulnerableto colonizationby alien floraand thenfunction sitesof as prolificseed production,vegetationmanagement programs shouldbe established thatintensively, but sensitively, managenaturereserveroadsides.Roadside vegetationmanagementprogramsshould (i) monitor statusof establishedalien species popthe ulations and the arrivalof new alien species, (ii) minimizeseed productionof keystonealien plant species-those having ecosystem-level effects (Macdonald et al. 1989), and (iii) rehabilitate roadsides usingeffective nativeseed mixesand substrate preparation.With the cooperation of local weed control agencies, managementprograms should also create buffer zones on roadsides leading into naturereserves. Even relatively simpleactions,such as mowingroadsidestowardrather thanaway from potential sourcesofalien seeds-for example,mowing outward to reserve boundaries-can be effective parts of cohesive roadside managementprograms. 4. Specifictreatments, including possible use ofherbicides, for keystonealien plant species should be establishedand approved prior to immigration of such species into naturereserves.Quick and decisiveproactivepolicies can yieldextremely effective results,even over large areas, as in the control of Centaurea maculosa in the province of Alberta, Canada (Ali 1989). 5. Because hay and manuremaycontainseeds of alien species, livestock-related activitiesshould be carefullyreevaluated.To allow seeds to be expelled prior to use on trails,effective quarantineprocedures should be established.Elimination private of in livestock naturereservesshouldbe seriously contemplated.

Acknowledgments
We are grateful Glacier NationalPark'sresearchand to resource management staff who supported this study, especiallyKathy MarDimont,Carl Key,Dave Lang,Cliff and Frank tinka, JackPotter, RachelPotter. Forcella,Carl Key, Marcel Rejmanek,and two anonymousreviewers reviewedearlierdrafts thismanuscript were very of and a helpfulin clarifying number of our ideas. We also thank for advice and AndyTyAndyMatchett statistical ser forassisting with fieldwork. Financialsupportfor thisstudy was providedbyGlacierNationalParkand the of University Wyoming-National Park Research Center,Laramie, Wyoming.

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Literature Cited Ali,S. 1989. Knapweed eradication programin Alberta. Pages 105-106 in P. K Fay and J.R. Lacey,editors.Proceedingsof theknapweedsymposium, April4-5, 1989. Extension Service, MontanaStateUniversity, Bozeman,Montana,EB 45. Anderson, C., K T. Harper,and R. C. Holmgren.1982. FacD. the of soil torsinfluencing development cryptogamic crustsin Utah deserts.Journal Range Management of 35:180-185. Baker,H. G. 1986. Patterns plantinvasionin North of America. Pages 44-57 in H. A. Mooneyand J.A. Drake,editors.Ecology ofbiologicalinvasionsofNorthAmericaand Hawaii.SpringerVerlag,New York. of Bazzaz, F. A. 1986. Lifehistory colonizingplants:some defeatures. mographic, genetic,and physiological Pages 96-110 in H. A. Mooney and J.A. Drake,editors.Ecology ofbiological invasionsof NorthAmericaand Hawaii. Springer-Verlag, New York. Bright, A. 1986. Hiker impact on herbaceous vegetation J. along trailsin an evergreenwoodland of centralTexas. Biological Conservation 36:53-69. of Brotherson, D., and S. R. Rushforth. J. 1983. Influence crypof togamic crusts on moisturerelationships soils in Navajo NationalMonument, Arizona.GreatBasinNaturalist 43:73-78. in Butt, J.Lacey,and G. Kennett.1989. Variation biomassof C., and three spottedknapweedunder threelevels of defoliation levels ofcompetition. Pages 95-99 in P. K Fayand J.R. Lacey, editors.Proceedingsof the knapweed symposium, April4-5, 1989. ExtensionService,MontanaStateUniversity, Bozeman, Montana,EB 45. D. R. Caldwell,M. M.,J.H. Richards, A.Johnson, S. Nowak,and R. S. Dzurek. 1981. Copingwithherbivory: caphotosynthetic pacity and resource allocation in two semiaridAgropyron bunchgrasses. Oecologia 50:14-24. of Cole, D. N. 1987. Effects three seasons of experimental on and communities a grassland trampling fivemontaneforest in western Montana,USA. Biological Conservation40:219244. and R. L. Iman. 1981. Ranktransformationsa as Conover,W. J., bridge between parametric and nonparametricstatistics. AmericanStatistician 35:124-129. N. Cronquist, A. H. Holmgren, H. Holmgren, L. Reveal,and A., J. P. K Holmgren.1977. Intermountain flora.Volume 6, Monocotyledons.Columbia University Press,New York. effects vegetation on Dale, D., and T. Weaver. 1974. Trampling ofthe trailcorridors northRockyMountainforests. of Journal ofApplied Ecology 11:767-772. WashR. of Daubenmire, 1970. Steppevegetation Washington. StationTechnical Bulletin ingtonAgricultural Experimental 62:1-131. A. Finklin, I. 1986. A climatichandbook for Glacier National Park-with data forWaterton Lakes NationalPark.USDA Forest ServiceIntermountain ResearchStationGeneralTechnical ReportINT-204,Ogden, Utah.

Forcella, and S.J.Harvey.1983. Eurasian F., weed infestation in westernMontana in relationto vegetationand disturbance. Madrono 30:102-109. Frenkel, E. 1970. Ruderalvegetationalong some California R. roadsides.University California of Publicationsin Geography 20:1-163. Hall, C. N., and F. R. Kuss. 1989. Vegetationalterationalong trailsin ShenandoahNational Park,Virginia. Biological Conservation 48:211-227. Hitchcock,C. L., and A. Cronquist.1973. Flora of the Pacific Northwest. University Washington of Press,Seattle,Washington. Koterba, D., andJ.R. Habeck. 1971. Grasslands theNorth W. of Forkvalley,GlacierNationalPark,Montana.CanadianJournal of Botany49:1627-1636. Lacey, C. A., J.R. Lacey, T. K Chicoine, P. K Fay, and R.A. French. 1986. Controlling knapweed on Montana rangeland. Circular311. Cooperative ExtensionService,Montana State University, Bozeman,Montana. Locken,L.J.,and R. G. Kelsey. 1987. Cnicinconcentrations in Centaureamaculosa spottedknapweed.BiochemicalSystematics and Ecology 15:313-320. Macdonald,I. A.W., L. L. Loope, M. B. Usher,and 0. Hamann. 1989. Wildlifeconservationand the invasion of nature reservesby introduced species: a globalperspective. Pages 215255 in J.A. Drake, H. A. Mooney,F. di Castri,et al., editors. Biologicalinvasions. Wiley& Sons,New York. Mack,R. N. 1981. InvasionofBromus tectorumL. into western NorthAmerica:an ecological chronicle.Agro-Ecosystems 7:145-165. Mack,R. N. 1986. Alienplantinvasioninto the Intermountain West:a case history. Pages 191-213 in H. A. Mooney and J.A. Drake,editors.EcologyofbiologicalinvasionsofNorthAmerica and Hawaii. Springer-Verlag, York. New Mack, R. N. 1989. Temperate grasslandsvulnerableto plant invasions: characteristics consequences. Pages 155-179 in and J.A. Drake,H. A. Mooney,F. di Castri, al. editors.Biological et invasions. Wiley & Sons,New York. Mack, R. N., and J.N. Thompson. 1982. Evolution in steppe with few large, hooved mammals. American Naturalist 119:757-773. Mooney,H. A.,S. P. Hamburg, J.A. Drake. 1986. The invaand sions of plantsand animalsinto California. Pages 250-272 in H. A. Mooney and J.A. Drake, editors.Ecology of biological invasionsof NorthAmericaand Hawaii. Springer-Verlag, New York. W. Mueggler, F., and W. L. Stewart.1980. Grasslandand shrubland habitattypesof westernMontana.USDA ForestService GeneralTechnical ReportINT-66,Ogden, Utah. Park Records. 1939. Correspondencebetween D. S. Libbey, Glacier ParkSuperintendent, G. W. Noffsinger, and President of the Park Saddle Horse Company,dated August 14, 1939. Archives. GlacierNationalPark,West Glacier,Montana.

Conservation Biology Volume 6, No. 2, June 1992

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Alien Flora Adjacent RoadandTrail to Corridors

& Tyser Worley Tyser, W.,and C. H. Key. 1988. Spottedknapweedin natural R. area fescue grasslands:an ecological assessment.Northwest Science 62:151-160. A. Watson, K, and A.J.Renney.1974. The biologyofCanadian weeds. 6. Centaureadiffusaand C. maculosa CanadianJournal of PlantScience 54:687-701. Weaver,T., and B. Woods. 1985. The exotic floraof Glacier National Park: a preliminary atlas. MSU Biology Report 37. MontanaStateUniversity, Bozeman,Montana. Weaver,T., and B. Woods. 1986. The exotic floraof Grand Teton NationalPark:an environmental atlas.MSU Report 38. MontanaStateUniversity, Bozeman,Montana. L. Wilkinson, 1987. SYSTAT.Evanston, Illinois. M. Williamson, 1989. Mathematical models of invasion.Pages A. et 329-350 inJ. Drake,H. A. Mooney,F. di Castri, al. editors. Biologicalinvasions. Wiley & Sons,New York. Williamson, and K C. Brown.1986. The analysisand modM., elling of Britishinvasions.PhilosophicalTransactionsof the RoyalSocietyof London 314:505-522. Wilson,S. D. 1989. The suppressionof nativeprairieby alien species introducedfor revegetation.Landscape and Urban Planning 17:113-119. Young,J.A.,R A. Evans,andJ.Major. 1972. Alienplantsin the GreatBasin.Joumalof Range Management 25:194-201.

by fixation algae Reddy,G. B., and J.Giddens. 1975. Nitrogen soil on fescuegrass crusts.Soil Science SocietyofAmericaProceedings 39:654-656. Pages of M. Rejmanek, 1989. Invasibility plant communities. 369-388 in J.A. Drake, H. A. Mooney,F. di Castri,et al., editors.Biological invasions.Wiley & Sons,New York. NaRuhle,G. C. 1972. Roads and trailsof Waterton-Glacier Minnesota. Minneapolis, J. tionalParks. W. Forney, D. Saunders, A., R.J.Hobbs, and C. R. Margules.1991. Biologa ical consequences of ecosystemfragmentation:review.Conservation Biology5:18-32. Schmidt, W. 1989. Plant dispersal by motor cars. Vegetatio 80:147-152. wear tolerR. Shearman, C., and J.B. Beard. 1975. Turfgrass and anamorphological, ance mechanisms:III. Physiological, wear tolerassociated with turfgrass tomical characteristics ance. Agronomy Journal 67:215-218. in P. Stringer, W. 1973. An ecological studyofgrasslands Banff, of CanadianJournal LakesNationalParks. and Jasper, Waterton Botany51:383-411. in R. Tyser, W. 1991. Ecologyof fescuegrasslands GlacierNaof tional Park. Final Report. University Wyoming-National Wyoming. ParkServiceResearchCenter,Laramie, Tyser,R.W. 1992. Vegetationassociatedwithtwo alien plant Montana. in species in a fescuegrassland GlacierNationalPark, GreatBasin Naturalist In press. 52.

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